Thresholds in the Species–Area–Habitat Model: Evidence from the Bryophytes on Continental Islands

Aim: To clarify whether (1) there are thresholds in the species–area–habitat relationship for bryophytes and potential mechanisms, (2) such thresholds vary among different bryophyte groups, and (3) choros is better than area or habitat alone in the prediction of SR. Location: Islands in central and southern Zhejiang, China. Methods: We investigated the species richness (SR) of five bryophyte groups (total bryophytes, total mosses, liverworts, acrocarpous mosses, and pleurocarpous mosses) and habitat types on 66 islands. By using four threshold models, the logarithmic and the power models, we quantified their SR–choros relationships (SKRs), species–area relationships (SARs), and species–habitat relationships (SHRs). We also conducted path analyses to detect the direct effects of area per se and habitat per se on the SR. Results: The AICc values of the SKR models were overall smaller than those of the respective SAR and SHR models. The left-horizontal two-threshold model was best for the SKRs. A phenomenon (the small-choros effect, SCE) in which SR independently varied choros below a given threshold was detected. The SCE thresholds were smaller in mosses than in liverworts and in acrocarpous mosses than in pleurocarpous mosses. No direct and positive effects of habitat per se on the SR were detected below choros thresholds for all five groups. Main conclusions: There were two thresholds and SCEs in the SKRs of all five bryophyte groups. The SCEs likely resulted from the elimination of the direct and positive effects of habitat diversity on the SR of the bryophytes on small choros islands. The SCE thresholds were high for species groups sensitive to environments. Choros was better than area or habitat alone in determining the SR of the bryophytes on continental islands.